An ordinal classification for te families of flowering plants

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Volume 85
Number 4

1998

Annals
of the
Missouri
Botanical
Garden

AN ORDINAL

CLASSIFICATION FOR THE
FAMILIES OF FLOWERING
PLANTS

The Angiosperm Phylogeny Group

Abstract

Recent cladistie analyses are revealing the phylogeny of flowering plants in increasing detail, and there is support
(or the monophyly of many major groups above the family level. With many elements of the major branch i rig sequence
oi phylogeny established, a revised suprafamilial classification of flowering plants becomes both feasible and desirable.
Here we present a classification of 462 flowering plant families in 40 putatively monophyletie orders and a small
number of monophyletie, informal higher groups, The latter are the monoeots, commelinoids, eudieots, core eudicots,
rosids including eurosids I and Q, and asterids including euasterids I and IL Under these informal groups there are
also listed a number of families w r ithout assignment to order. At the end of the system is an additional list of families
of uncertain position for which no firm data exist regarding placement anywhere within the system.

Why rearrange families, still less formalize or-
ders? Higher-level classifications, the grouping of
species into families, orders, etc., are needed as
reference tools not only in systematics but also in
many other branches of biology. Knowledge of phy-
logenetic relationships of major groups of organ-
isms, that is, a phylogenetic perspective, is becom-
ing increasingly important, and hence the need for
a phylogenetic classification as a reference tool is
also becoming imperative.

Our primary focus is on orders with a secondary
emphasis on families of flowering plants. The family
is central in flowering plant systematics. For ex-
ample, in studying an unknown plant we usually
first identify it to family. The orders, on the other
hand, have until quite recently been of little im-
portance, either being morphologically unrecogniz-
able or in most cases lacking any evolutionary co-
herence (Heywood, 1977; Merxmiiller, 1977).
However, orders are useful in teaching, for studying

1 Recommended citation, abbreviated as "APG, 1998/* This paper was compiled by Ktfre Bremer, Mark W. Chase,
and Peter F Stevens, equally responsible and listed here in alphabetical order only, with contributions from Arne A.
Anderberg, Anders Baeklund, Birgitta Bremer, Barbara G« Briggs, Peter K. Endress, Michael F. Fay, Peter ColdhlaH,
Mais H. G. Gustafsson, Sara B. Hoot, Walter S. Judd, Mari Kiillersjo, Elizabeth A. Kellogg, Kathleen A. Kron, Donald
H* Les, Cynthia M, Morton, Daniel L. Nickrent, Richard G. Olmstead, Robert A. Price, Christopher J. Quiim, James
E, Rodman, Paula J. Rudall, Vincent Savolainen, Douglas E. Soltis, Pamela S. Soltis, Kenneth J. Sylsma, and Mats
Thulin (in alphabetical order). Addresses: K. Bremer, Department of Systematic Botany, Uppsala University, Villavagen
6, S-752 36 Uppsala, Sweden; M. W. Chase, Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9
3DS, U.K.; P. F. Stevens, Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A.

Ann. Missouri Bqt\ Card. 85: 531-553, 1998,

532

Annals of the
issouri Botanical Garden

family relationships, and in positioning genera of
doubtful affinity. The didactic value of suprafami-
lial groupings has been emphasized by various au-
thors (e.g., Dahlgren, 1975; Thorne, 1976; Davis,
1978; Takhtajan, 1997). This value is even more
evident now that the phylogeny of flowering plants
is being disclosed in increasing detail, Many of the
orders recognized by earlier authors are not mono-
phyletic, yet there is a pressing need for names to
communicate the knowledge of monophyletic
groupings of families that are becoming evident.
With the major branching sequence of flowering
plant phylogeny becoming clearer, a revised famil-
ial and ordinal classification is feasible.

Flowering plant classification systems from the
late 1970s seemed to be stable ami show substan-
tial agreement, but this stability has been rudely
shattered as new kinds of data and new methods of
analyzing conventional data have become firmly es-
tablished (Stevens, 1986). Classifications such as
those by Cronquist (1981) and Takhtajan (1980),
although still in frequent use, have become outdat-
ed. Of more recent classifications, that by Goldberg
(1986) of the dicotyledons predates the advent of
molecular studies at higher levels, as does that by
Dahlgren et al. (1985) of the monocotyledons. How-
ever, the latter incorporated much new data and
provided synapomorphy schemes for many groups.
The recent system of Takhtajan (1997), although
extremely elaborate, is made less useful because
his propensity for splitting often results in well-
known families being dismembered, then reassem-
bled as orders. Furthermore, the findings of recent
molecular studies, despite being cited, have hardly
influenced his classification.

We conclude that there is a great need for a new,
phylogenetic classification of flowering plants, pro-
viding names for major monophyletic groups of
families* Obviously, it is not possible, nor is it de-
sirable, to name all cladcs in the entire phylogeny.
Any such complete classification would be so cum-
bersome that it would be useless for general com-
munication. Systematists need to come to some
kind of agreement concerning which clades to rec-
ognize and name, so that a reference tool of broad
utility can be formulated and used to discuss di-
versity. An ordinal classification of flowering plant
families is here proposed for that purpose (pp. 538—
542), It recognizes a selected number of monophy-
letic suprafamilial groups, that is, clades in the
phylogeny of flowering plants that are supported by
at least one, and often several, lines of evidence.
These are clades to which we find it useful to refer
when we communicate information about higher-
level interrelationships of the flowering plants.

We note that the selection of clades to be rep-
resented in a formal classification is different from
the procedure of naming these clades* The latter
issue of biological nomenclature in phylogenetics is
currently much debated (e.g., Cantino et al., 1997;
de Queiroz, 1997; Lid£n et ah, 1997), but we have
not adopted any "phylogenetic naming" sensu de
Queiroz and Gauthier (1994). We operate under the
current International Code of Botanical Nomencla-
ture (Greuter et ah, 1994) and choose to emphasize
the ranks of family and order. The Linnaean cate-
gories serve as a convenient mnemonic device for
remembering hierarchical relationships, but it
should of course be realized that groups of the same
rank are evolutionarily non-comparable units un-
less they are sister groups.

There are noteworthy problems when establish-
ing the names for taxa at ordinal and other higher
taxonomic levels. Until recently, little attention has
been paid to the nomenclature at these levels, and
our knowledge of the early literature in which such
names were used is imperfect. This situation has in
considerable part been rectified by Reveals (1998)
Herculean labors. The principle of priority is not
mandatory for taxa above the rank of family, al-
though authors are exhorted "generally" to follow
this principle (Greuter et al., 1994). We have tried
to balance priority with general usage when assign-
ing names to orders, but even if future bibliograph-
ic work discloses earlier ordinal names, changes
are not mandated.

Which clades should be recognized in classifi-
cation, or in our case, how should the orders be
circumscribed? Given the primary principle of
monophyly, that of recognizing clades ami not
grades in classification, there are nevertheless
many considerations to be taken into account when
circumscribing taxa at ordinal as well as all other
hierarchical levels above that of species. Classifi-
cation is not only a matter of grouping according to
the principle of monophyly, but it is also a matter
of communication (note that whatever philosophy of
naming is adopted, there has to be some consensus
as to the clades we are going to use in general
botanical communication). For us, this raises the
question of ranking, that is, after having selected
clades in the phylogeny to be named, they have to
be assigned an appropriate place in the hierarchy,
in our case, family and order (e.g., Backlund & K,
Bremer, 1998; Stevens, 1998). In choosing between
alternative circumscriptions it is desirable to rec-
ognize groups that are well supported. It is also
useful to select groups that have some kind of eas-
ily observed morphological synapornorphies, al-
though this may be difficult at the ordinal level and

Volume 85, Number 4
1998

Angiosperm Phytogeny Group
Ordinal Classification

OJv

even sometimes at the family level. Synapomor-
phies also often include (sometimes exclusively)
anatomical, biochemical, and developmental char-
acters.

Many of our ordinal names are already well es-
tablished and used in earlier classifications and
systematic treatments. So far as they represent
monophyletic groups, we retain well-known orders
in the interest of preserving stability. In other eases,
the size of the orders comes into consideration.
However, what is reasonably broad circumscrip-
tion? From the point of view of memorization of
names, groups of 2—6 or a few more would seem to
be ideal, and there is evidence that systematists in
the past have commonly recognized groups of this
size (Stevens, 1997), However, with the discoveries
of new species, genera, and families, the sizes of
genera, families, and orders have increased* and
many orders now comprise 10-20 families, or even
more. Other orders contain a few families only, and
if there are only two or three families in an order,
"one is not far from leaving the families unplaced"
(Copeland, 1957). Concerns about the doubtful val-
ue of recognizing similarly small groups have also
been expressed by others (e.g., Burtt, 1977), Nev-
ertheless, we have chosen to recognize a number of
small orders because these represent clades for
which monophyly and relationships are well sup-
ported, and this better conveys the interrelation-
ships of the families included rather than leaving
them unclassified to order.

In general, we adopt a broad circumscription of
the orders* We recognize 462 families and 40 or-
ders of flowering plants. Cronquist (1981) recog-
nized 321 families and 64 orders, Thome (1992)
440 families and 69 orders, and Takhtajan (1997)
no less than 589 families in 232 orders. Our wider
ordinal circumscription is not because finer details
of the phylogeny within the orders are as yet un-
clear, but because we think the classification will
be more useful with a limited number of larger or-
ders. As we develop more firmly supported phytog-
enies within and among orders, groups at the in-
fraordinal and supraordinal levels can be
recognized. Hence we anticipate that there will be
little need to change the circumscription of the or-
ders recognized here, except for inclusion of yet
unassigned families of unknown systematic position
and the transfer of misplaced families. Additional
orders may have to be recognized as the phyloge-
netic relationships of families that are not yet
placed are clarified. Discussion as to whether a
widely accepted monophyletic group should be a
superorder, order, suborder, or family is largely vac-

uous because this will always be an arbitrary de-
cision.

Takhtajan (1997) opted in favor of "smaller, more
natural families and orders, which are more coher-
ent and better-defined, where characters are easily
grasped, and which are more suitable for informa-
tion retrieval and phylogenetic studies, including
cladistic analyses (e.g., because it reduces poly-
morphic codings)/* However, the size of a group has
nothing to do with its "naturalness." For a smaller
group, one will often be able to say more about all
of its constituent members, and so the characters
may be more easily grasped. However, segregates
of well established monophyletic families like Ru-
biaceae (Gentianales) or Asteraceae (Asterales)
would by Takhtajan s generalization also be more
natural; by this criterion, the smaller the group, the
more natural it will necessarily be, so there is no
ranking criterion to be derived from "naturalness."
If by "more natural" is meant "has more synapo-
morphies" then this, too, is incorrect; the number
of syuapomorphies is not connected to the size of
the group or the hierarchical level at which it is
recognized.

In our classification, these considerations have
had little impact. The principle of monophyly in
combination with the desirability of maintaining al-
ready well established and familiar entities has
largely formed the ordinal classification. Monofam-
ilial orders (and monogeneric families) are avoided
as much as possible, minimizing redundancy in
classification. In a few cases we have, however, rec-
ognized some monofamilial orders (Ceratophyllales,
Acorales, Arecales) because these are sister groups
of more than one other order. Hence, the families
of these monofamilial orders cannot be included in
any other order without violating monophyly.

The principle of monophyly in combination with
the mandatory usage of the family category (Greuter
et aL, 1994) may lead to the recognition of many
small families. For example, in Dipsacales, if Dip-
sacaceae and Valerianaceae are to be retained as
families separate from Caprifoliaceae, the principle
of monophyly requires the recognition also of Dier-
villaceae, Linnaeaceae, and Morinaceae (Backlund
& K, Bremer, 1998; Backlund & Pyck, 1998). This
is because each of these latter families is the sister
group of more than one family so they cannot be
merged with any other family without violating
monophyly. Similar considerations apply at the or-
dinal level. Unfortunately, no absolute guidelines
as to reasonable practice can be offered, but we
simply observe that caution is always in order.

In other cases there are small families that may
be reduced to synonymy of their sister group if the

534

Annals of the

Missouri Botanical Garden

latter consists of a single family. Examples are Ca-
bombaceae, which may be merged with Nymphae-
aceae, and Kingdoniaceae, which may be merged
with Gireaeasteraceae (Ranunculales), Such com-
monly recognized families that nevertheless may be
merged with their sister family are in our classifi-
cation placed within square brackets below the
family with which they may be merged (in Ran-
unculales either Fumariaceae or both Fumariaceae
and Pteridophyllaceae may be merged with Papav-
eraeeae; alternatively, either Pteridophyllaceae or
both Fumariaceae and Pteridophyllaceae may be
retained as distinct).

We do not attempt to thoroughly revise family
circumscriptions. In general we follow recent au-
thors and attempt to recognize as many monophy-
letic families as possible. It should be emphasized,
however, that following additional investigation
some families listed below may be shown to be non-
monophylctic; revised circumscriptions, either by
merging or splitting, into inonophy letic taxa are not
yet possible given our current knowledge. Exam-
ples are Euphorbiaceae and Flacourtiaceae of Mai-
pighiales (Kallersjo et ah, 1998) and several fami-
lies of Myrtales (Conti et aL, 1996; Gadek et al.,
1996) and core Caryophyllales (which comprise
Achatocarpaceae, Aizoaeeae, Amaranthaceae, Bas-
ellaeeae, Caetaceac, Caryophyllaceae, Didierea-
ccae, Molluginaceae, Nyctaginaceae, Phytolacca-
ceae, Portulaeaeeae, Sarcobataceae, and
Stegnospermataceae; Hershkovitz & Zimmer,
1997)* Other probably non-monophyletic families
that cannot yet be recircumscribed are Boragina-
ceae {euasterids I; Chase et al., 1993), Scrophular-
iaceae (Lamiales; Olmstead & Reeves, 1995), and
Santalaceae (San tal ales; Nickrent & Duff, 1996;
Nickrent et aL, 1998). Brassicaceae (Brassicales)
include also the former, paraphyletic Capparaeeae
(Brassicaceae sensu stricto being nested inside
Capparaceae; Judd et aL, 1994; Rodman et aL,
1996). A supposedly parallel rase comprises Api-
aceae and Araliaceae (Apiales), since the former
have been assumed to be nested inside the latter
(Plunkett et aL, 1996). However, with a transfer of
Hydrocotyloideae from Apiaceae to Araliaceae, it
seems that two monophyletic families can be rec-
ognized, only a few genera remaining unplaced
(Plunkett et aL, 1997). Delimitation of Bombaca-
ceae, Malvaceae, Sterculiaceae, and Tiliaceae
(Mai vales) is problematical, and only Malvaceae
are monophyletic (Alverson et al M 1998; Bayer et
aL, 1999). Here all four are treated together as a
single monophyletic family, Malvaceae sensu lato
(Judd & Manchester, 1997),

Our proposed classification is a modification of

that conceived by Bremer et al. (1995, 1996, 1997)
and since 1996 available on the Internet (Bremer
et aL, 1998). This classification is based on various
recently published mostly molecular phylogenetic
analyses (e.g., Chase et aL, 1993; Chase et aL,
1995; Bremer et al,, 1994; Struwe et aL, 1994; Na-
dot et aL, 1995; Nickrent & Soltis, 1995; Soltis et
aL, 1995; Gadek et aL, 1996; Gustafsson et aL,
1996; Morton et aL, 1996; Soltis & Soltis, 1997;
Soltis et aL, 1997; Anderberg et aL, 1998; Back-
lund & B. Bremer, 1998; Bakker et aL, 1998; Kal-
lersjo et aL, 1998; Soltis et al., 1998; Thulin et al.,
1998; further references above). The major differ-
ences are in the expansion of Alismatales (includ-
ing also Araceae), Caryophyllales (including Dro-
seraceae, Nepcnthaceae, Polygonaceae, Plumbagi-
naeeae, and several other families outside the tra-
ditional, core Caryophyllales), the recognition of a
comparatively widely circumscribed Rosales (in-
cluding Rhamnaceae, Urticaceae, Moraceae, and
their allies), in the addition of a number of smaller
orders (Ceratophyllales, Acorales, Arecales, Prote-
ales, Garryales, Aquifoliales), and in the deletion
of a few r others (Aristoloehiales, Nymphaeales, Bro-
meliales, Trochodendrales, Zygophyllales), Mono-
cots and eudicots are not formally ranked and
named because it is not yet clear at which level
they should be recognized. The same problems oc-
cur with commelinoids (a phylogenetically derived
subgroup of monocots) and with rosids and asterids
(subgroups of eudicots), although these are com-
monly known as subclasses Commelinidae, Rosi-
dae, and Asteridae, respectively.

Well supported ordinal interrelationships are
shown in Figure 1. Interrelationships among the
basal branches of the tree and the position of the
root of the flowering plant phylogeny remain elu-
sive. Within the eudicots there is increasing sup-
port for a large subgroup with predominantly pen-
tamerous and isomerous flowers, the core eudicots,
mainly comprising Caryophyllales, Santalales, Sax-
ifragales, rosids, and asterids. Rosids and asterids
each comprise two large subgroups, eurosids I and
II and euasterids I and II, also receiving increasing
support as monophyletic. These correspond to the
similarly numbered rosid and asterid clades of
Chase et al. (1993).

Under each of the supraordinal groups of mono-
cots, commelinoids, core eudicots, rosids, etc.,
there are a number of families listed without as-
signment to order These families are known to be-
long within the major group under which they are
listed, but their ordinal position is still uncertain.
Similarly, Amborellaceae, Austrobaileyaceae, Ca-
nellaceae, etc., are listed at the beginning because

Volume 85, Number 4
1998

Angiosperm Phytogeny Group
Ordinal Classification

535

1M/76

100/54

Ceratophyllales
Lau rales
Magnoliales
Piperales

100/100

0>
CO

o

■a

CD

3
c/>

3
o

o
o

99/55

50/58

90/68

CO

96/100

84/87

100/99

08/-

07/-

-/-

/-

82/-

Acorales
Alismatales

Asparagales

Dioscoreales

Liliales

Pandanales

Arecales

Poaies

Commelinales

J222L Zingiberales

commelinoids

100/60

CD

O

o

CO

58/-

100/99

100/98

100/-

o

CD
Q.

o"

O

99/58

O
0)

77/55

68/-

-/-

60/

CO

Ranunculales

Proteales

Caryophyllales

Santalales

Saxifragales

Geraniales

Malpighiales

Oxali dales

Fabales

Rosales

Cucurbitales

Fagales

95/

si

00/72

00/-

06/77

00/94

00/80

eurosids I

00/90

00/94

00/50

00/98

88/-

100/-

C/>
CD

CO

W/68

88/-

100/89

.

100/86

N/-

mm

00/-

87/-

100/82

H/8S

88/98

Brassicales

Malvales

Sapindales

Cornales

Ericales

Garryales

Gentianales

Lamiales

Solanales

Aquifoiiales

Apiales

Aste rales

Dipsacales

eurosids II

euasterids I

euasterids It

Figure 1. Phylogenetic interrelationships of the orders of (lowering plants, compiled from recent cladistir analyses
cited in the text. J ark knife support is given on the branchrs (a {lash for values < 50^), first jaekknife values from
analysis of 545 sequences of the r//rL, atpli m and 18S rDNA genes (D. E. Sollis, M. W. Chase, R S. Soltis, D. Albach,
M. E. Mort, V. Saiolainen, M. Zanis & J* S, Karris, unpublished, in prep.) and second jackkuife values from analysis
of 2538 rbd, sequences (Kallersjo e-t aL 1998).

536

Annals of the

Missouri Botanical Garden

they belong neither in any of the phylogeiietically
"basal*' orders at the beginning nor in the monorots
or eudicots. Furthermore, families listed directly
under monorots without an order are monorots but
not rommelinoids, and families similarly listed di-
rectly under eudicots and core eudieots are eudi-
cots or core eudicots, respectively, but neither ros-
ids nor asterids. At the end of the system is an
additional list of families of uncertain position.
Most of these are probably eudicots (including core
eudicots* rosids, and asterids), but so far there are
no firm data supporting their placement anywhere
within the eudicots.

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of characters. Bot. Not. 128: 119-147.

, IL T, Clifford & P. F. Yen. 1985, The Families of

the Monocotyledons. Spri tiger- Verlag, Berlin.

Davis, P. H. 1978. The moving staircase: An analysis of
taxouomic rank and affinity* Notes Roy. Bot. Card. Ed-
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De (Jueiroz. K> |W7, The Linnaean hierarchy and the
evolutioni/ation of taxonomy, with enmhasis on the
prcmlem of nomenclature. Aliso 15: 125-144,

& J. Gauthier. 1W4. Toward a phylogenetic sys-
tem of biological nomenclature. Trends EcoL Evol. 9:
27-31,

(iadek, P. A.. E. S. Fernando, C. J. Quinn, S. B. Hoot. T.
Terrazas, VI. C* Sheahan &' M. W". Chase. 19*Xx Sap-
indales: Molecular delimitation and infraordinal groups.
Amer. J. Bot. 83: 802-81 L

. P. <;. Wilson & C. J. Quinn, 1996. Phylogenetic

reconstruction in Myrlaccae u>ingmnfK, with particular
reference to the position of Psiloxyfon and Heteropyxis.
Austral. Svst. Bot. 9: 283-290.

Coldberg. A. |9W>. Classification, evolution, and phvlog-
env of the farm lies ol dicotvled<»ns. Smithsonian Contr.
Hot. 58: 1-314.

(rreuter, W., K K. Barrie, FL VL Burdet. \\. (i. Chaloner,
V, Hcmouliii. I). L. HawkswortlK P. M. Ji*rgensen, 1). IL
Nicolson. P. C. Silva, P. Trehane & J. McNeill. 1994.
International Code ol Botanical Nomenclature. Hegnuni
Veg. 131.

(iustafsson, M. H. G., A. Backlund & B, Bremer. 1996.
Phvlogen\ of the Asterales sensu lato based on rhcL
sequences with particular reference to ihe Coodeni-
aceae, PL Syst. Evol. 1W: 217-242.

Hershkovilz. M. A. & E. A. Zimtner. IW7. On the evo-
lutionary origins of the cacti, laxon 46: 217-232.

Heywood, \. H. 1977. Principles and concepts in the {'las-
si heat ion of higher laxa. In K. Kuhitzki (editor). Flow-
ering plants: Evolution and classification oi higher cat-
egories. Pi. Syst, EvoL Suppl. 1: 1-12.

JudtL Vt. S, & S + R. Manchester. 1997. Circumscription

Volume 85, Number 4
1998

Angiosperm Phylogeny Group
Ordinal Classification

537

of Malvaceae (Mai vales) as determined by a preliminary
eladistic analysis of morphological, anatomical, paly-
nological, and chemical characters. Brittonia 49: 384—
405.

• R. W. Sanders & M, J, Donoghue. 1994, Angio-

sperm family pairs: Prelim inary phylogenetic analyses.
Harvard Pap. Bot. 5: 1-51.

K&llersjfr, M., J. S. Farris„ M. W, Chase, B. Bremen M. F.
Pay, C. J. Humphries. C. Petersen, (). Scbrrg & K.
Bremer. 1998. Simultaneous parsimony jaekknife anal-
ysis of 2538 rhcL DNA sequences reveals support for
major chides of green plants, land plants, seed plants,
and flowering plants. PI. SysL Evol. [in press].

I j den, M., B. Oxelman, A. Baeklund, L. Andersson, B.
Bremer. R. Eriksson, R. Moherg, 1. Nordah K. Persson,
M. Thulin & B, Zimmer. 1 W7. Charlie is our darling.
Taxon 46: 73 .>- 738,

Merxmiiller, H. 1977. Summarv lecture. In K. Kubttzki
(editor)* Flowering plants: Evolution and classification
of higher categories. PL Syst, Evol. Suppl. 1: 397—105.

Morton, C. M., M, W. Chase, K. A. Kron & S, M, Swensen.
1996, A molecular evaluation of the monophvly of the
order Ebenales based upon r/jrL sequence data. Svst.
Bot. 2 i : 567-586.

Nadot, S., G. Bittar. L, Carter, R. Lacroix & B. Lejeune.
1995, A phylogenetic analysis of monocotyledons based
on the chloroplast gene rps4 using parsimony and a new
numerical phenetics method. Molee. Phvlog. Evol. 4:

257-282.

Nickrent, D. L & J. R. Duff. 1996, Molecular studies of
parasitic plants using ribosomal RNA. Pp. 28-52 in M.
T, Moreno, J. L Cubero, I). Rerner, D. Joel, L J. Mus-
selman & C. Parker (editors). Advances in Parasitic
Plant Research. Junta de Andalucfa, Direccion General
de lnvesiigaci6n Agraria, Ctfrdoba, Spain.

& I). E. Soltis. 1995. \ comparison of angiosperm

phylogenies from nuclear 18S rDNA and rhrL sequenc-
es. Ann. Missouri Bot. Gard. 82: 208-234.

, J. R. Duff, A. E. CoIwelL A. D. Wolfe, N. D.

Young, k, K. Steiner & C. W. dePamphilis, 1998. Mo-
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sitic plants. Pp. 211-241 in D. E. Sohis, P. S. Soltis &
J, J. Doyle (editors), Molecular Systematic* of Plants II:
DNA Sequencing. Kluwer, Boston.

Olmstead. R. C + & P. A. Reeves. 1995. Evidence for the
polyphyly of the Scrophulariaceae based on chloroplast
r6cL and ndhY sequences. Ann. Missouri Bot. Gard. 82:
176-193.

Plunkett, G, M„ D. E. Soltis ft P. S. Soltis. 1996, Higher
level relationships of Apiales (Apiaceae and Araliaceae)
based on phylogenetic analysis of rhrl, sequences.
Amen J. Bot, a3: 399-415.

. & . 1997. Classification of the

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matK and r&cL sequence data, Amer. J. Bot. JJ4: 56;
580.

Reveal, J. L. 1998. Indices nominum supragenericorum
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edu/PBIOA WWW7supragen.html.

Rodman, J. E., K. G. Karoh R. A. Price & K. J. Sytsnia.
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Soltis, D. E. & P. S. Soltis. 1997. Phylogenetic relation-
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Amer, J. Rot. 84: 504-522.

. . 1). R. Morgan, S. M. Swensen, B. C.

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, . D. L. Nickrenl, L. A. Johnson. \fc. J.

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1-49.

Stevens. R F. 1986, Evolutionary classification in botany,
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Thome, R, F. 1976, A phylogenetic classification of the
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Thulin, M., B. Bremer, J. Richardson, J, Niklasson, M, F,
Fay & M. W, Chase, 1998, Family relationships of the
enigmatic rosid genera Barhexa and Dirarhma from the
Horn of Africa region. PL Syst. Evol, 213: 103-119.

538

Annals of the

Missouri Botanical Garden

Classification ok Flowkrim; Plants

Arnborellaceae

Austrohaileyaeeae

Canellaceae

Chloranthaceae

Hydnorareae

Illiciaceae

Nymphaeaceae

[ + Cabombaceae]

Rafflesiaeeae

Schisandraceae

Triineniaceae

Winteraceae

Ceratophy Hales Bisch,
Ceratophyllaeeae

Laurales Perleb
Atherosperniataceae
Calyc ant haceae
Gornortegaceae
Hemandiaerae
Lauraeeae
Monimiaceae
Siparunaeear

Magnoliales Bromhead
Annonaceae
Degeneriaceae
Eupomatiaceae
Himantandraceae
Magnoliaceae
Myristicaceae

Piperales Dumort.
Aristolochiaceae
Lac tori darea**
Piperaceae
Saururaceae

MONOCOTS

Corsiaceae

Japonoliriaceae

Nartheciaceae

Petrosaviaceae

Triuridaceae

A co rales Reveal
Acoraceae

Alismatales Dumort,
Alisriiataceae

Araceae

Butomaceae
Cymodoceaceae

Hy droc har i taceae
Juncaginaceae

Limn oehari taceae

Posidoniaeeae

Potamogetonaceae

Ruppiaceae

St 1 heucfa zeri ac e ae

Tofieldiareae

Zosteraeeae

Asparagales Bromhead
Agapanthaceae
Agavaceae
Alliaceae
Amaryllidaceae
Anemarrhenaceae
Anthericaceae
Aphyllanthaceae
Asparagaeeae
Asphodelaceae
Asteliaceae
Behniaceae
Blandfordiaceae
Boryaceae
Convallariaceae
Doty ant haceae
Hemeroeallidaceae
Herreriaceae
Hesperocallidaceae
Hyacint haceae
Hypoxidaceae

Iridaceae
Ixioliriaeeae

Lanariaceae
Laxmanniaceae
Orchidaceae
Tecophilaeaceae

Tlieinidureae

Xanthorrhoeaceae

Xeroneniataceae

Dioscoreales Hook* L
Burmanniarcae
Dioscoreaceae
Taccaceae

Thismiaceae
Trirhopodaceae

Li Hales Perleb
Alstroemeriaceae
Campynemataceae
Colchieaeeae
Liliaceae
Luzuriagaeeae
Melanthiaceae
Philesiaceae
Ripogonaceae
Smilacaceae

Volume 85, Number 4
1998

Angiosperm Phytogeny Group
Ordinal Classification

Doy

Classification of Flowering Plants
(cont'd,)

Pandanales LindL
Cyclanthaceae
Pandanaceae

Stemonaeeae
Velloziaceae

COMMELINOIDS

Abolbodaceae

Bromeliaceae

Dasy pogo n ac e ae

Hanguanaceae

Mayacaceae

Rapateaceae

Arecales Bromhead
Arecaceae

Commelinales DumorL
Commelinaceae
Haemodoraceae

Philydraeeae
Pontederiaceae

Poales Small

Anarthriaceae
Centrolepidaceae
Cyperaceae
Ecdeiocoleaceae

Eriocaulaceae

Flagellariaceae

Hydatellaceae

Joinvilleaceae

Juncaceae

Poaceae

Prioniaceae

Restionaceae

Sparganiaeeae

Thurniaeeae

Typhaceae

Xyridaceae

Zingiberales Griseb.
Cannaceae
Costaceae
Heliconiaceae
Lowiaceae
Marantaceae
Musaceae
Strelitziaceae
Zingiberaceae

EUDICOTS

Buxaceae

Didymelaceae

Sabiaceae

Troehodendraceae
[ +Tetracentraceae]

Proteales Dumort.
Nelumbonaceae
Platanaceae
Proteaceae

Ranunculales Dumort,
Berberidaceae
Circaeasteraceae
[ + Kingdoniaceae]
Eupteleaceae
Lardizabalaceae
Menispermaceae
Papaveraeeae
[ + Fumariaeeae]
[ + Pteridophyllaceae]
Ranunculaceae

CORE EUDICOTS

Aextoxicaeeae

Berberidopsidaceae

Dilleniaceae

Gunneraceae

Myrothamnaceae

Vitaceae

Caryophyllales Perleb
Achatocarpaceae
Aizoaceae
A niarant haceae
A nc i strocladaeeae
Asteropeiaceae
Basellaceae
Cactaceae
Caryophyilaceae
Didiereaceae
Dioncophyllaeeae
Droseraceae
Drosophyllaceae
Frankeniaceae
Molluginaceae
Nepenthaceae
Nyctaginaceae
Physenaceae
Ph y t o 1 ac c ac eae
Plumbaginaceae
Polygonaceae
Portulacaceae
Rhabdodendraceae
Sarcobataeeae
Simmondsiaceae
Stegnospermataceae
Tamarieaeeae

Santalales Dumort.
Olacaceae
Opiliaoeae

540

Annals of the

Missouri Botanical Garden

Classification ok Flowkrim; Plants
(cont'd.)

Loranthaceae

Misodendrateae

Santalaeear

Saxifragales Dumort,
Altingiaceae
Cerridiphyllaceae
Crassulaceae
Daphniphyllareae
Grossulariaceae
Haloragaeeae
Hamamelidaceae
Iteaceae

Paeoniaceae

Penthoraceae
Pterostemonaceae
Saxifragaceae
Tetrararpaeaceae

ROSIDS

Aphlniareae

Crossosoinatareae

Ixerbaoeae

Krameriaceae

Pirranitiiareae

Podostetnarrat 1

Staehyuraceae
Staph yleaceae
Tristichaceae
Zygophyllaceae

Geraniales Dumort,
Francoaccae
Geraniaceae
[ + Hypseofharitareae]
Greyiaeeae
Ledoearpaceae
Melianthaceae
Vivianiaceae

EUROSIDS I

Celastraceae

Huaceae

Parnassiaeeae

[ + Lepuropetalaceae]

Staekhousiaeeae

Cueurbitales Dumort.

Anisophylleaceae

Rrgnniaceae

Coriariareae

Corynoearpaeeae

Cucurbitaceae

Datiscaceae

IVtramelaceae

Fabales Rromhead
Fabaceae
Polygalactia**
Quillajaceae
Surianaceae

Fagalcs Engl.
Retulateae
Casuarinaceae
Fagaceae
Juglandaceae
Myricaceae
Nothofagaceae
Rhoipteleaceae
Ti eodend rac eae

Malpighiales Mart.
Achariaceae
Ralanupaceae
Garyocaraceae
Chrysobalanareae
Clusiaceae
Di chape talaceae
Erythroxylaceae
Euphorbiaeeae
Euphroniaceae
Flacourtiaceae
Goupiaceae
1 lugnniaccae
Humiriaceae
Irvingiaceae
Ixonanthaoeae
Lacistemataeeae
Linaceae
Malesherbiaceae
Malpighiaccae
Mcdusagynaceae
Gchnaceae
Pandaceae
Passifloraccae
Putranjivaceae
Quiinaceae
Rhizophoraceae
Salicaceae
Scyphostegiaccac
Trigoniaceae
Tumeraceae
Violaceae

Oxalidales Heintze
Cephalotaceae
Connaraceae
Cunoniaceae
Elacocarpaeeac
Oxalidaceae
Tremandraceae

■

Volume 85, Number 4
1998

Angiosperm Phylogeny Group
Ordinal Classification

541

Classification of Flowering Plants
(cont'd-)

Rosales Perleb
Barbeyaceae
Cannabaceae
Cecropiaceae
Celtidaceae
Dirachmaceae
Elaeagnaceae
Moraceae
Rhamnaceae
Rosaeeae
Ulmaceae
Urticaceae

EUROSIDS II
Tapisciaceae

Brassicales Bromhead
Akaniaceae
[+Bretsehneideraceae]
Bataceae
Brassicaeeae
Caricaceae
Emblingiaceae
Gy ro s t e monat ■ eae
Koeberliniaceae
Limnanthaceae
Moringaceae
Pentadiplandraceae
Resedaceae
Salvadoraceae
Setchellanthaceae
Tovariaceae
Tropaeolaceae

Mai vales Dumort.
Bixaeeae

[ + Diegodendraceae]
Cistaceae
Cochlospermaceae
D i pteroc arpac eae
Malvaceae
Muntingiaceae
Neuradaceae
Sareolaenaceae
Sphaerosepalaceae
Thymelaeaceae

Myrtales Rchb*
Alzateaceae
Combretaceae
Crypteroniaceae
Heteropyxidaceae
Lythraceae
M e 1 as tomatac eae
Memecylaceae
Myrtaceae
Oliniaceae

Onagraceae

Penaeaceae

Psiloxylaceae

Rhy nc hoc aJ y c aceae

Voehysiaeeae

Sapindales Dumort.
Anaeardiaceae
Biebersteiniaceae
Burseraceae
Kirkiaceae
M el i aceae
Nitrari aceae
[ + Peganaceae]
Rutaceae
Sapindaceae
Simaroubaceae

ASTERIDS

Cornales Dumort.

Cornaceae

[H-Nyssaceae]

Grubbiaceae

Hydrangeaceae

Hydrost achy aceae

Loasaceae

Ericales Dumort.
Actinidiaceae
Balsaminaceae
Clethraceae
Cyrillaceae
Diapensiaceae
Ebenaceae
Ericaceae
Fouquieriaceae
Halesiaceae
Lecythidaceae
Maregraviaceae
Myrsinaceae
Pellicieraceae
Pol emoni aceae
Primulaceae
Roridulaceae
Sapotaceae
Sarraceniaceae
Styracaceae
Symplocaceae
Tenistroemiaceae
Tetrameristaceae
Theaceae
Theophrastaceae

EUASTERIDS I

Boraginaceae
Plocospermataceae

Vahliaceae

542

Annals of the

Missouri Botanical Garden

Classification of Flowering Plants

(cont'd)

Garry ales Lindl.
Aucubaceae
Eucommiaceae
Garryaceae
Oncothecaceae

Gentianales Lindl.
Apocynaceae
Gelsemiaceae
Gentianaceae
Logan iaceae
Rubiaceae

Lamiales Bromhead
Acanthaceac
Avicenniaceae
Bignon iaceae
Buddlejaceae
Byblidaceae
Cyclocheilaecae
Gesneriaceae
Lam iaceae
Lentibulariaceae
Myoporaceae
Oleaceae
Orobanehaceae
Paulowniaceae
Pedaliaeeae
[ + Marty niaceae]
Phrymaceae
Plantaginaeeae
Schlegel iaceae
Serophulariaceae
Stilbaceae
Tetraehondraceae
Verbenaceae

Solanales DumorL
Convolvulaceae
Hydroleaceae
Monti niaceae
Solanaceae
Sphenocleaceae

EUASTERIDS II

Adoxaceae

Bmniaceae

Carlematmiaceae

Columclliaceae

[ + Desiontainiaceaej

E r e mosy n ae eae

Escalloniaceae

Icacinaceae

Polyosmaceae

Sphenostemoriaeeae

Tribclaeeae

Apiales Nakai
Apiaceae
Araliaceae
Aralidiaeeae
Griseliniaeeae
Melanophyllaceae
Pittosporaceae
Torricelliaceae

Aquifoliales Senft
Aquifoliaceae
Helwingiaceae

Phyllonomaceae

Asterales Lindl.
Alseuosmiaceae
Argophyllaceae
Asteraceae
Calyceraceac
Gainpanulaceae
[+Lobeliaeeae]
Carp ode taceae
Donatiaceae
Goodeniaceae
Menyanthaceae
Pentaphragmataceae
Phellinaceae
Rousseaceac
Stylidiaceae

Dipsacales DumorL
Capnfoliaceac
Diervillaceae
Dipsaeaeeae
Linnaeaceae
Morinaceac
Valerianaceae

Families of Uncertain Position

Balanophoraceae

Bonnetiaceae

Cardiopteridaceae

Ctenolophonaceae

Cynomoriaceae

Cytinaeeae

Dipentodontaceae

Elatinaceac

Geissolomataceae

Hoplest i gmataceae

Kaliphoraceae

Lepidobotryaceae

Lissocarpaceae

Lophopyxidaceae

Medusandraceae

Metteniusaceae

Mitrastenionaceae

Paracryphiaceae

Volume 85, Number 4
1998

Angiosperm Phytogeny Group
Ordinal Classification

543

Classification or Flowkring Plants

(cont*d.)

Pentaphylacaceae

Peridiscaceae

Plagiopteraceae

Pottingeriaceae

Sladeniaceae

Strasburgeriaceae

Tepuianthaceae

Ordinal Synonyms

Acanthales Lindl.

= Lamiales
Acerales Lindl.

= Sapindales
Actinidiales Takht. ex Reveal

= Ericales
Adoxales Nakai

- not accepted, family under

euasterids II
Aesculales Bromhead

= Sapindales
Agavales Hutch,

= Asparagales
Alliales Traub

= Asparagales
Alstroemeriales Hutch,

= Liliales
Altingiales Doweld

= Saxifragales
Amaranthales Dumort.

= Caryophyllales
Amaryllidales Bromhead

= Asparagales
Ambrosiales Duniort.

= Asterales
Am mi ales Small

= Apiales
Amomales Lindl.

= Zingibe rales
Ancistrocladales TakhL

= Caryophyllales
Annonales Lindl.

= Magnoliales
Anthobolales Dumort.

= Santalales
Apocynales Bromhead

= Gentianales
Aponogetonales Hutch.

= Alismatales
Arales Dumort.

= Alismatales
A rali ales Reveal

= Apiales
Aralidiales Takht. ex Reveal

= Apiales
Aristolochiales Dumort.

= Piperales

Asarales Horan.

= Piperales
Asclepiadales Dumort.

= Gentianales
Asteliales Dumort.

= Asparagales
Atriplicales Horan*

= Caryophyllales
Aucubales Takht.

= Garryales
Austrobaileyales Takht. ex Reveal

- not accepted, family at beginning
of system

Avenales Bromhead

= Poales
Balanopales Engl,

= Malpighiales
Balanophorales Dumort.

- not accepted, family unplaced
Balsaminales LindL

= Ericales
Barbeyales Takht, & Reveal

= Rosales
Batales Engl.

= Brassicales
Begoniales Dumort.

= Cucurhitales
Berberidales Dumort.

= Ranunculales
Brtu lales Bromhead

= Fagales
Biebersteiniales Takht.

= Sapindales
Bignoniales LindL

= Lamiales
Bixales Lindl.

= Malvales
Boraginales Dumort.

- not accepted, family under
euasterids I

Brexiales LindL

- not accepted, family under
eurosids I

Bromeliales Dumort.

- not accepted, family under
commelinoids

Bruniales Dumort.

- not accepted, family under
euasterids II

Brunoniales LindL

= Asterales
Burmanniales Heintze

= Dioscoreales
Burserales Baskerville

= Sapindales
Butomales Hutch.

= Alismatales

544

Annals of the

Missouri Botanical Garden

Classification nv Flovvkkim; Plants
(cont'd*)

Buxales Takht. ex Reveal

- not accepted, family under
eudieots

Byblidales Nakai ex Reveal

= Lamiales
Gactales Duinort

= Caryophyllales
Callitriehales Dumort,

= Lamiales
Calyeanthales Mart.

= Lau rales
Calyccralcs Takht, ex Reveal

= Asterales
Gampanulales Rchb,

= Asterales
Canellales Cronquist

- not accepted, family at beginning
of system

Cannales Dumort.

= Zingiberales
Capparales Hutch.

= Brassicales
Caprifoliales Lindl.

= Dipsaeales
Cardiopteridales Tak ht .

- not accepted, family under
euasterids II

Carduales Small

= Asterales
Caricales L. D. Benson

= Brass ic ales
Cassiales Horan.

= Fabales
Casuarinales Lindl.

= Fagales
Celastrales Baskerville

- not accepted, family under
eurosids I

Centrolepidales Takht.

= Poales
Cephalotales Nakai

= Oxalidales
Cercidiphyllales Hu ex Reveal

= Saxifragales
Chenopodiales Dumoit

= Caryophyllales
Chiruniales Griseb.

= Gentianales
Chloranthales A. C. Sm. ex J. -F, Leroy

- not accepted, family at beginning
of system

Cinchonales Lindl.

= Gentianales
Circaeasterales Takht,

= Ranunculales
Cistales Rchb.

= Mai vales

Citrales Duniort.

= Sapindales
Cocosales Nakai

= Arecales
Colchicales Duniort.

= Liliales
Comhretales Baskerville

= Myrtales
Connarales Takht. ex Reveal

= Cunoniales
Convolvulales Dumort.

= Solanales
Coriariales Lindl.

= Cueurbitales
Corylales Dumort.

- Fagales
Corynocarpales Takht.

= Cueurbitales
Crassulales LindL

= Saxifragales
Crossosornatales Takht. ex Reveal

- not accepted, family under rosids
Cunoniales Hutch,

= Oxalidales
Cyclanthales J. 1L Schaffn,

= Pandanalcs
Cvniodoeealcs Nakai

= Alisma tales
Cynomoriales Burnett

- not accepted, family unplaced
Cyperales Hutch,

= Poales
Cytinales Dumort.

- not accepted, family unplaced
Daphnales Lindl.

= Malvales
Daphniphyllales Pulle ex Cronquist

= Saxifragales
Datiseales Dumort,

= Cueurbitales
Desfontainiales Takht.

- not accepted, family under
euasterids II

Diapensiales Engl, & Gilg

= Ericales
Didymelalcs Takht.

- not act epted, family under
eudieots

Dillcniales Hutch.

- not accepted, family under core
eudieots

Dioneophy Hales Takht. ex Reveal

= Gary ophy Hales
Diospy rales Prantl

= Ericales
Droserales Griseb,

= Caryophyllales

Volume 85, Number 4
1998

Angiosperm Phylogeny Group
Ordinal Classification

545

Classification of Flowering Plants
(cont'd.)

Ebenales Engl.

= Ericales
Elaeagnales Bromhead

= Resales
Elaencarpales Takht.

= Oxalidales
Elatinales Nakai

- not accepted, family unplaced
Elodeales Nakai

= Alismatales
Empetrales Raf.

= Ericales
Eriocaulales Nakai

= Poales
Eucommiales Neinejc ex Cronquist

= Ganyales
Euphorbiales Lindl.

= Malpighiales
Eupomatiales Takht, ex Reveal

= Magnoliales
Eupteleales Hu ex Reveal

= Ranunculales
Euryalales H.L.Li

- not accepted, family at beginning
of system

Ficales Dumort.

= Rosales
Flacourtiales Heintze

= Malpighiales
Fouquieriales Takht, ex Reveal

= Ericales
Francoales Takht.

= Geraniales
Frangulales Wirtg.

= Rosales
Galiales Bromhead

= Gentianales
Geissolomatales Takht. ex Reveal

- not accepted, family unplaced
Gesneriales Dumort.

~ Lamiales
Glaucidiales Takht. ex Reveal

- Ranunculales
Globulariales Dumort.

= Lamiales
Goodeniales Lindl.

= Asterales
Greyiales Takht.

= Geraniales
Grossulariales LindL

= Saxifragales
Gunnerales Takht. ex Reveal

- not accepted, family under core
eudicots

Gyrocarpales Dumort.
= Laurales

Gyros temonales Takht.

= Brassicales
Haemodorales Hutch,

= Commelinales
Haloragales Bromhead

= Saxifragales
Hamamelidales Griseb.

= Saxifragales
Hanguanales R. Dahlgren ex Reveal

= not accepted, family under

commelinoids
Helleborales Nakai

= Ranunculales
Helwingiales Takht.

= Aquifoliales
Himantandrales Do weld & Shevyryova

= Magnoliales
Hippuridales Pulle ex Reveal

= Lamiales
Homaliales Bromhead

= Malpighiales
Hortensiales Griseb,

= Cornales
Hydatellales Cronquist

= Poales
Hydnorales Takht. ex Reveal

- not accepted, family at beginning
of system

Hydrangeales Nakai

= Cornales
Hydrastidales Takht.

= Ranunculales
Hydropeltidales (BartL) Spenn.

- not accepted, family
Nymphaeaceae at beginning of
system

Hydrostachyales Diels ex Reveal

= Cornales
Hypcricales Dumort.

- Malpighiales
Hypoxidales Takht.

= Asparagales
Icacinales Tiegh. ex Reveal

- not accepted, family under
euasterids II

Illiciales Hu ex Cronquist

- not accepted, family at beginning
of system

Iridales Raf.

= Asparagales
Ixiales LindL

= Asparagales
Jasminales Dumort.

= Lamiales
Juglandales Dumort.

= Fagales

546

Annals of the

Missouri Botanical Garden

Classification ok Flowkrim; Plants
(cont'd*)

Julianialcs Engl.

= Sapindales
Juneaginales Hutch,

= Alismatales
Juncales Dumort.

= Poales
Lacistematales Baskerville

= Malpighiales
Lac tori dales Takht, ex Reveal

= Piprrales

Lardizabalales Loconte

= Ranunculales
Lecythidales Cronquist

= Ericales
Leitneriales Engl.

= Sapindales
Lentibulariales Lindl.

= Lamiales
Ligustrales Bartl. ex Bisch.

= Lamiales
Limnanthales Nakai

= Brassirales
Linales Baskerville

= Malpighiales
Loasales Bessey

= Cornales
Loganiales Lindl.

= Gentianales
Lonicerales T. Liebe

= Dipsacales
Loranthales Duinort.

= Santalales
Lythrales Caruel

= Mvrtales
Marat hrales Dumort.

- not accepted, family
Podostemaceae under rosids

Mayacales Nakai

- not accepted, family under
commelinoids

Medusagynales Takht.

= Malpighiales
Medusandrales Brenan

- not accepted, family unplaced
Melanthiales R, Dahlgren ex Reveal

= Liliales
Melastoniatales Oliv,

= Myrtales
Meliales LindL

= Sapindales
Menispermales Bromhead

= Ranunculales
Menyanthales T. Yamaz. ex Takht,

= Asterales
Metteniusales Takht,

- not accepted, family unplaced

Mitrastemonales Makino

- not accepted^ family unplaced
Monimialcs Dumort.

= Lau rales
Moringales Nakai

= Brassicales
Myricales Engl.

= Fagales
Myristicales Thom6

= Mag noli ales
Myrothamnales Nakai ex Reveal

- not accepted, family under core
eudieots

Myrsinales Spenn.

= Ericales
Najadales Dumort.

= Alismatales
Narcissales Dumort.

= Asparagales
Nartheciales Reveal & Zomlefer

- not accepted, family under
monocots

Nelumbonales Reveal

= Proteales
Ne pent hales Dumort.

= Caryophyllales
Nolanales Lindl.

= Solanales
Nyctaginales Dumort.

= Caryophyllales
Nymphaeales Dumort.

= not accepted, family at beginning

of system
Ochnales Hutch* ex Reveal

= Malpighiales
Qenotherales Bromhead

= Myrtales
Olacales Benth.

= Santalales
Oleales Lindl.

= Lamiales
Onagrales Rchb,

= Myrtales
Opuntiales Willk.

= Caryophyllales
Orchidales Raf.

= Asparagales
Paeoniales Heintze

= Saxifragales
Pandales Engl. & Gilg

= Malpighiales
Papaverales Dumort.

= Ranunculales
Paracryphiales Takht.

- not accepted, family unplaced
Paridales Dumort.

= Liliales

Volume 85, Number 4
1998

Angiosperm Phytogeny Group
Ordinal Classification

547

Classification of Flowering Plants

(cont'd.)

Parnassiales Nakai

- not accepted, family under
eurosids I

Passifiorales DumorL

= Malpighiales
Penaeales LindL

= Myrtales
Petiverialcs LindL

= Caryophyllales
Petrosaviales Takht.

- not accepted, family under
monocots

Phi lyd rales DumorL

= Commelinales
Physenales Takht

= Caryophyllales
Pinguiculales Dumort.

= Lamiales
Pittosporales LindL

= Apiales
Plantaginales LindL

= Lamiales
Platanales J. H. Schaffn.

= Proteales
Plumbaginales LindL

- Caryophyllales
Podophyllales Dumort.

= Ranunculales
Podostemales LindL

= not accepted, family under rosids
Polemoniales Bromhead

= Ericales
Polygalales Dumort,

= Fabales
Polygonal es Dumort.

= Caryophyllales
Pontederiales Hook, f.

= Commelinales
Portulacales Dumort.

= Caryophyllales
Posidoniales Nakai

= Alismatales
Potamogetonales Dumort,

= Alismatales
Primulales Dumort.

= Ericales
Quercales Burnett

= Fagales
Raff Irs tales 01 i v.

- not accepted, family at beginning

of system
Resedales Dumort.

= Brassicales
Restionales J. H, Schaffn.

= Poales
Rhamnales Dumort.

= Rosales

Rhinanthales Dumort.

= Lamiales
Rhizophorales Tiegh. ex Reveal

= Malpighiales
Rhodorales Horan.

= Ericales
Rhoipteleales Novdk ex Reveal

= Fagales
Roridulales Nakai

= Ericales
Rubiales Dumort.

= Gentianales
Ruppiales Nakai

= Alismatales
Ru tales Perleb

= Sapindales
Sabiales Takht.

= not accepted, family under

eudicots
Salicales LindL

= Malpighiales
Salvadorales R, Dahlgren ex Reveal

= Brassicales
Samolales Dumort*

= Ericales
Samydales Dumort.

= Malpighiales
Sanguisorbales Dumort.

= Rosales
Sapotales Hook. f.

= Ericales
Sarraceniales Bromhead

= Ericales
Scheuchzeriales B. Boivin

= Alismatales
Sclerant hales Dumort.

= Caryophyllales
Scrophulariales LindL

= Lamiales
Scyphostegiales Croizat

= Malpighiales
Sedales Rchb.

= Saxifragales
Silenales LindL

= Caryophyllales
Simmondsiales Reveal

= Caryophyllales
Smilacales LindL

= Liliales
Stellariales Dumort,

= Caryophyllales
Stylidiales Takht* ex Reveal

= Asterales
Sty rac ales Bisch.

- Ericales
Taeeales Dumort.

= Dioscoreales

548

Annals of the

Missouri Botanical Garden

Classification ok Flowekino Plants

(cont'd.)

Tamales DuniorL

= Dioseoreales
Tamaricales Hutch.

= Caryophyllales
Tecophilaeales Traub ex Reveal

= Asparagales
Theales Lindl.

= Ericales
Theligonales Nakai

= Gentianales
Thymelaeales Willk.

= Malvales
Tiliales Camel

= Malvales
Tofieldiales Reveal & Zomlefer

= Alismatalrs
Torricelliales Takht. ex Reveal

= Apiales
Tovariales Nakai

= Brassieales
Trilliales Takht.

= Liliales
Triuri dales Hook. f.

- not accepted, family under monocots
Trochodendrales Takht. ex Cronquist

- not accepted, family under
eudicots

Tropaeolales TakhL ex Reveal

= Brassieales
Turnerales Dumort,

= Malpighiales
Typhales Dumort.

= Poales
Ulmales Lindl.

= Rosales
Urticales Dumort.

= Rosales
Vacciniales Dumort.

= Ericales
Vallisneriales Nakai

= Alismatales
Velloziales R. Dahlgren ex Reveal

= Pandanales
Veratrales Dumort.

= Liliales
Verbenales Horan,

= Lamiales
Viburnales Dumort.

- not accepted, family under
euasterids II

Vincales Horan.

88 Gentianales
Violates Perleb

= Mai pigh Sales
Vitales Reveal

- not accepted, family under core
eudicots

Voehysiales Dumort.

= Myrtales
Winterales A. C, Sm. ex Reveal

- not accepted, family at beginning
of system

Xyridales Lindl.

= Poales
Zosterales Nakai

= Alismatales
Zygophyllales Takht.

- not accepted, family under rosids

Selected Familial Synonyms

Abrophyllaceae

= Carpodetaceae
Acanthoehlamydaceae

= Velloziaceae
Aceraceae

= Sapindaecae
Achradaceae

= Sapotaceae
Aegicerataeeae

= Myrsinaceae
Agdestidaeeae

= Phytolaceaeeae
Aitoniaceae

= Meliaceae
Alangiaceae

= Cornaceae
Aloaceae

= Asphodelaceae
Alsinaceae

= Caryophyllaceae
Ambrosiaceae

= Asteraceae
Amygdalaceae

= Rosaceae
Audrostachyaceae

= Euphorbiaceae
Antoniaceae

= Logan iaceae
Apodanthaceae

= Raffles iaceae
Apostasiaceae

= Orchidaceae
Aptandraceae

= Olacaceae
Aristoteliaceae

= Elaeocarpaceae
Asclepiadaceae

= Apocynaceae
Asteranthaceae

= Lecythidaceae
Averrhoaceae

= Oxalidaceae
Avetraceae

= Dioscoreaeeae

Volume 85, Number 4
1998

Angiosperm Phytogeny Group
Ordinal Classification

549

Classification of Flowering Plants
(cont'd,)

Balanitaceae

= Zygophyllaceae
Barbeiuaceae

= Phytolaecaceae
Barclayaceae

= Nymphaeaceae
Barri ngtoniaceae

= Lecythidaceae
Baueraceae

= Cunoniaceae
Baxteriaceae

= Dasypogonaceae
Bembici aceae

= Flacourti aceae
Berzeliaeeae

= Bruniaceae
Bisehofiaeeae

= Euphorbiaceae
Blepharocaryaceae

= Anacardiaceae
Boerlagellaceae

= Sapotaceae
Bombacaceae

= Malvaceae
Boopidaceae

= Calyceraceae
Bretschneideraceae

= Akaniaceae
Brexiaceae

= Celastraceae
Brunelliaceae

= Cunoniaceae
Brunoniaeeae

= Goodeniaceae
Bumeliaceae

= Sapotaceae
Burchardiaceae

= Colehicaceae
Byttneriaceae

= Malvaceae
Cabombaceae

= Nymphaeaceae
Caesalpiniaceae

= Fabaceae
Calectasiaceae

= Dasypogonaceae
Callitrichaceae

= Plantaginaceae
Calochortaceae

= Liliaceae
Camelliaceae

= Theaceae
Canotiaceae

= Celastraceae
Cansjeraceae

= Opiliaceae

Capparaceae

= Brassicaceae
Carduaceae

= Asteraceae
Cassythaceae

= Lauraceae
Chailletiaceae

= Dichapetalaceae
C henopodi aceae

= Amaranthaceae
Chionographidaceae

~ Melanthiaceae
Chloanthaceae

= Lamiaceae
Cichoriaceae

= Asteraceae
Cleomaceae

= Brassicaceae
Cneoraceae

= Rutaceae
Cobaeaceae

— Polemoniaceae
Compositae

= Asteraceae
C onosty 1 i dac eae

= Haemodoraceae
Cordiaceae

= Boraginaceae
Corid aceae

= Primulaceae
Corokiaceae

= Argophyllaceae
Corylaceae

= Betulaceae
Croomi aceae

= Stemonaceae
Cruciferae

= Brassicaceae
Curtisiaceae

= Comaceae
Cuseutaceae

= Convolvulaceae
Cyananthaceae

= Campanulaceae
C yan as tr aceae

= Tecophilaeaceae
Cynocrambaceae nom. illeg.

= Rubiaceae
Cyphiaceae

= Campanulaceae
C y phoc arpac eae

= Campanulaceae
Cypripediaceae

= Orchidaceae
Dactylanthaceae

= Balanophoraceae

550

Annals of the

Missouri Botanical Garden

Classification ok Flowering Plants
(cont'd,)

Davidiaceae

= Cornaeeae
Davidsoniaceae

— Cunoniaeeae
Deeaisneaceae

= Lardizabalaceae
Desfontainiaceae

= Columelliaceae
D i aly petal an thaeeae

= Rubiaceae
Dianellaceae

= Hemerocallidaceae
Dichondraceae

= Convolvulaceae
Diclidantheraeeae

= Polygalaeeae
Dirgodendrareae

= Bixaeeae
Dionaeaeeae

= Droseraceae
Dracaenaceae

— Convallariaceae
Duabangaceae

= Lythraeeae
Duckeodendraceae

= Solanaieae
Dulongiaceae nom. illeg.

= Phyllonomaceae
Dysphaniaceae

= Amaranthaceae
Ehretiaceae

= Boraginaceae
EllLsiophyllaceae

= Scrophulariaceae
Empetraeeae

= Ericaceae
Epacridaeeae

= Ericaceae
Eremolepidaceae

= Santalaceae
Eriospermaeeae

= Convallariaceae
Erycibaceae

= Convolvulaceae
Erythropalareae

= Olacaceae
Eucryphiaceae

= Cunoniaceae
Euryalaceae

= Nvmphaeaceae
Exoearpaceae

= Santalaceae
Flindersiaceae

= Rutaceae
Foetidiaceae

= Lecythidaceae

Frangulaceae

= Rhamnaeeae
Fumariaeeae

= Papaveraceae
Funkiaceae

= Agavaceae
Galacaceae

= Diapensiaceae
Geitonoplesiaceae

= Hemerocallidaceae
Geniostomaceae

= Loganiaceae
Geosiridaceae

= Iridaeeae
Gisekiaceae

= Phytolaccaceae
Glaueidiaeeae

= Ranunculaccae
Globulariaceae

= Plantaginaeeae
Goetzeaceae

= Solanaceae
Gonystylaeeae

= Thymelaeaceae
Gouaniaceae

= Rhamnaeeae
Gramineae

= Poaceae
Gronoviaceae

= Loasaeeae
Gustaviaceae

= Lecythidaceae
Guttiferae

= Clusiaceae
Gyrocarpaceae

= Hernandiaceae
Halophilaceae

— Hydrocharitaceae
Halophytaceae

= Amaranthaceae
Hectorellaceae

= Portulacaceae
Heliotropiaceae

= Boraginaceae
Heloniadaceae

= Melanthiaecae

Helosidaceae

= Balanophoraeeae
Henriqueziaeeae

= Rubiaceae
Hippoeastanaceae

= Sapindaceae
Hippocrateaceae

= Celastraceae
Hippuridaceae

= Plantaginaeeae

Volume 85, Number 4
1998

Angiosperm Phylogeny Group
Ordinal Classification

551

Classification of Flowering Plants

(cont'd.)

Hortoniaeeae

= Monimiaceae
Hostaceae

= Agavaceae
Humbertiaceae

= Convolvulaeeae
Hydrastidaceae

= Ranunculaceae
Hydrocotylaceae

= Amliaceae
Hydropeltidaceae

= Nymphaeaceae
Hydrophyllaceae

= Boraginaceae
Hymenocardiaceae

= Euphorbiaceae
Hypecoaceae

= Papaveraeeae
Hypericaceae

= Clusiaceae
Hypseocharitaceae

= Geraniaceae
Idiospermaceae

= Calyeanthaceae
Illecebraeeae

= Caryophyllaceae
Jasionaceae

= Campanulaceae
Jasminiaceae

— Oleaeeae
Johnsoniaceae

= Hemerocallidaceae
Julianiaceae

= Anac ardiaceae
Kiggelariaceae

= Flaeourtiaceae
Kingdoniaceae

= Circaeasteraeeae
Kirengeshomaceae

= Hydrangeaceae
Labiatae

— Lamiaceae
Langsdorffiaceae

= Balanophoraceae
Leeaceae

= Vitaceae
Legurninosae

= Fabaceae
Leitneriaceae

= Simaroubaceae
Lemnaceae

= Araceae
Lennoaceae

= Boraginaceae
Leoniaceae

= Violaceae

Lepuropetalaceae

= Parnassiaceae
Lilaeaceae

= Juncaginaceae
Limoniaceae

= Plumbaginaceae
Liriodendraceae

= Magnoliaceae
Lobeliaceae

= Campanulaceae
Lomandraceae

= Laxmanniaceae
Lophiraeeae

= Ochnaceae
Luphophytaceae

= Balanophoraceae
Luxemburgiaceae

— Ochnaceae
Malaceae

= Rosaceae
Martyniaceae

= Pedaliaceae
Mastixiaceae

= Cornaceae
Medeolaceae

= Liliaceae
Meliosmaceae

= Sabiaceae
Mcndonciaceae

= Acanthaceae
Mesembryanthemaceae

= Aizoaceae
Mirnosaceae

= Fabaceae
Monotaceae

— Dipierocarpaeeae
Monotropaeeae

= Ericaceae
Mouririaceae

= Memecylaceae
Moutabeaceae

= Polygalaceae
Myriophyllaceae

= Haloragaceae
Mystropetalaceae

= Balanophoraceae
Najadaceae

= Hydrocharitaceae
Nandinaceae

= Berberidaceae
Napoleonaceae

= Lecythidaceae
Naucleaceae

= Rubiaceae
Nectaropetalaceae

= Erythroxylaceae

552

Annals of the

Missouri Botanical Garden

Classification ok Flowering Plants

(eontM.)

Nelson iaceae

= Acanthaeeae
N e m ac 1 ad aceae

= Campanulaceae
Nesogenaceae

— Cyclocheilaceae
Nolanaceae

= Solanaceae
Nolinaceae

= Convallariaceae
Nupharaeeae

= Nymphaeaceae
Nyetanthaceae

= Oleaeeae
Nyssaceae

= Cornaceae
Oetoknemaceae

= Olac aceae
Oft iaceae

= Scrophulariaeeae
Oph iopogonaeeae

= Convallariaceae
Osyridaeeae

= Santalaceae
Pachysandraeeae

= Buxaceae

Pal mat*

— Arecaceae
Papilionaceae

= Fabaceae
Peganaceae

= Nitrariaceae
Pentastemonaceae

= Stemonaceae
Peperomiaceae

= Piperaeeae
Periplocaceae

= Apocynaceae
Peripterygiaceae

= Cardiopteridaceae
Peterm an n iaceae

— Colehieaceae
Petiveriaceae

= Phytolaccaceae
Pliiladelphaeeae

= Hydrangeaceae
Phormiaceae

= Hemerocallidaceae
Phylicaceae

= Rhamnaceae
Picrodendraeeae

= Euphorbiaceae
Pinguiculaceae

= Lentibulariaceae
Pi stac iaceae

= A nacard iaceae

Pist iaceae

= Araceae
Platystemonaceae

— Papaveraceae
PI umer iaceae

= Apocynaceae
Podoaceae

— Anacardiaceae
Podophyllaceae

= Berberidaceae
Polygonanthaeeae

= Anisophylleaceae
Potal iaceae

= Gentianaceae
Ptaeroxylaceae

= Rutaceae
Pteridophyllaceae

= Papaveraceae
Punicaceae

= Lythraceae
Pyrolaceae

= Ericaceae
Ranzaniaceae

= Berberidaceae
Reaumuri aceae

= Tamaricaceae
Retz iaceae

= Stilbaceae
Rhinanthaceae

= Qrobanehaceae
Rhodoleiaceae

= Hamamelidaceae
Rhopalocarpaceae

= Sphaerosepalaceae
Rhynchothecaceae

= Ledoearpaccae
Roxburghiaceae

= Stemonaceae
Ruscaceae

= Convallariaceae
Saccifoliaceae

= Gentianaceae
Salaciaceae

= Celastraeeae
Salicorniaceae

= Amaranthaceae
Salpiglossidaceac

= Solanaceae
Sambucaceae

= Adoxaceae
Samolaceae

= Primulaceae
Sanicu Iaceae

= Apiaceae
Sarcophytaceae

— Balanophoraeeae

Volume 85, Number 4
1993

Angiosperm Phytogeny Group
Ordinal Classification

553

Classification of Flowebing Plants
(cont'd,)

Sarcospermataceae

= Sapotaceae
Sar ge n t od oxac eae

= Lardizabalaceae
Saurauiaceae

= Actinidiaceae
Sauvagesiaceae

= Qehnaceae
Scaevolaceae

= Goodeniaceae

~ Euphorbiaceae
Schoepfiaceae

= Olaeaeeae
Sclerophylacaceae

= Solanaeeae
Scoliopaceae

= Liliaceae
Scybaliaeeae

= Balanophoraceae
Scytopetalaceae

= Lecythidaceae
Selaginaceae

= Scrophulariaceae
Sesamaceae

= Pedaliaceae
Sesuviaceae

= Aizoaeeae
Sinietlndaceae

= Hemerocallidaceae
Siphonodontaceae

= Celastraeeae
Sonnerat iaceae

= Lythraceae
Spigeliaceae

= Loganiaceae
Stenomeridaceae

= Dioscoreaceae
Sterculiaceae

= Malvaceae
Stilaginaceae

= Euphorbiaceae
Strychnaceae

= Loganiaceae
Stylobasiaceae

= Surianaeeae
Stylocerataceae

= Buxaceae
Symphoremataceae

= Lamiaceae
Syringaceae

= Oleaceae
Tetracentraceae

= Trochodendraceae

Tetradiclidaceae

— Peganaceae
Tetragon iaceae

= Aizoaeeae
Tetrastylidiaceae

= Olacaceae
Thalassiaceae

= Hydroeharitaceae
Theligonaceae

= Rubiaceae
Thunbergiaceae

= Acanthaeeae
Tiliaceae

= Malvaceae
Trap ac eae

= Lythraceae
Trapellaceae

= Pedaliaceae
Tribu Iaceae

= Zygophyllaceae
Tricyrtidaceae

= Liliaeeae
Trilliaceae

= Melanthiaceae
Triplostegiaceae

= Valerianaceae
Uapacaceae

= Euphorbiaceae
Ullucaceae

= Basellaceae
Umbelliferae

= Apiaceae
Utrieulariaceae

= Lentibulariaceae
Uvulariaceae

= Colchicaceae
Vaccin iaceae

= Ericaceae
Vibumaeeae

— Adoxaceae
Viscaceae

= Santalaceae
Viticaceae

= Lamiaceae
Walleriaceae

= Tecophilaeaeeac
Wellstediaceae

= Boraginaeeae
Xanthophyllaceae

= Polygalaceae
Xerophyllaceae

= Melanthiaceae
Zann ichell i aceae

= Potamogetonaceae